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River Floodplains As Habitat and Bio-Corridors for Distribution of Land Snails: Their Past and Present

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River Floodplains As Habitat and Bio-Corridors for Distribution of Land Snails: Their Past and Present 10.1515/jlecol-2015-0012 Journal of Landscape Ecology (2015), Vol: 8 / No. 3. RIVER FLOODPLAINS AS HABITAT AND BIO-CORRIDORS FOR DISTRIBUTION OF LAND SNAILS: THEIR PAST AND PRESENT JITKA HORÁČKOVÁ1,2, ŠTĚPÁNKA PODROUŽKOVÁ1 AND LUCIE JUŘIČKOVÁ1 1Department of Zoology, Charles University in Prague, Faculty of Science, Viničná 7, CZ-128 44 Prague 2, e-mails: [email protected], stepanka.podrouzkova@ gmail.com, [email protected] 2Center for Theoretical Study, Charles University in Prague and the Academy of Sciences of the Czech Republic, Jilská 1, CZ-110 00 Prague 1 Received: 23rd October 2015, Accepted: 27th November 2015 ABSTRACT River floodplains of Czech rivers serve as refugia to woodland or hydrophilous gastropods,in current intensively agriculturally utilised, urbanised and largely fragmented landscape. This habitat often form one of the last refuge and replace the natural habitat of these species. River floodplains also represent linear bio-corridors in landscape and allow gastropods to spread through the landscape in both directions, up and down the stream. We showed based on available fossil mollusc successions that development of the floodplain mollusc fauna took place quite different way in various river floodplains, depending on their specifics and geographical location, because especially the ones situated in the chernozem area of the Czech Republic had very different history in comparison with those in higher altitudes. The species richness and composition of recent floodplain malacofauna arises from historical development of particular area/site and depends also on environmental factors such as an elevation, humidity gradient, vegetation type and its biomass, light conditions of the site and soil reaction. Recently, the invasive plants represent a serious problem for current floodplain ecosystems; species richness and abundances of terrestrial mollusc floodplain assemblages are changing due to their effect. The impact on gastropods is species-specific and was described for the following species: Impatiens glandulifera, Fallopia japonica subsp. japonica, F. sachalinensis, F. ×bohemica. Key words: terrestrial gastropods, river floodplain, bio-corridor, plant invasions, historic development INTRODUCTION River floodplain can be defined as a flat bottom of the valley with a characteristic biota, formed and influenced by the river stream (Štěrba et al., 2008; Ložek, 2007; 2011). River floodplains are one of the most dynamic ecosystems in Central Europe. They fulfil numerous functions in the landscape and their ecological research is therefore of a long tradition (Gurnell, 1977; Schnitzler et al. 2005), concerning to study of vegetation (Schnitzler, 1994; Horáčková J., Podroužková Š., Juřičková L. : River floodplains as habitat and bio-corridors for distribution of land snails: their past and present AAAAAAAAAAAAAAAAAAAAAAAAAAAAAAAAAAAAAAAAAAAAAAAA Brown et al. 1997). A number of regional studies have been carried out in the area of Central Europe since the second half of the 20th century (e.g. Ložek, 1955; Bába, 1977; Frank, 1984, 1985; Obrdlík et al., 1995; Čejka, 1999; 2003; Čejka et al. 2008; Ilg et al. 2009; Čejka & Hamerlík, 2009), and part of them were dealed with malacofauna of Czech rivers (e.g. Ložek, 1947; Horsák, 2000; Vašátko et al. 2002; Myšák & Horáčková, 2011; Horáčková et al. 2011a,b; 2013a,b,c; 2014a). Some studies were investigated the inundation influence on the river floodplain malacofauna (Ilg et al., 2009) or the function of river as bio-corridor (Myšák & Horsák, 2011) but the majority of the above mentioned studies is only descriptive and is missing the analysis of the main patterns of the species composition and diversity of floodplain mollusc assemblages in relation to ecological gradients of the environment. A large number of studies dealing with analysis of the main factors of the environment influencing the species diversity and composition of gastropod assemblages in various types of European temporal woodlands have been published (e.g. Wäreborn, 1969; Bishop, 1980; Martin & Sommer, 2004a; Cameron & Pokryszko, 2005; Pokryszko & Cameron, 2005; Hylander et al., 2005). However until now, the fauna of alluvial forests was neglected. Studies on the floodplain forests of the Danube River (Čejka et al., 2008) and the Elbe River including its tributaries (Horáčková et al., 2014b) were published in the last couple of years. Floodplain forests and riparian vegetation in the Czech river and brook floodplains serve as refugia for woodland and hydrophilous gastropods, because in the current fragmented and extensively human-utilized country are often ones of the last and hardly replaceable natural habitats. HISTORY OF RIVER FLOODPLAIN FORMATION If we are trying to understand the dynamics of river floodplains, we have to analyse them from the point of view of their establishment. Recently, there are 16 strictly river floodplain sites with fossil mollusc successions (Horáčková et al., 2015) within the area of the Czech Republic and Slovakia. They can help to understand the development of floodplain mollusc assemblages in the post-glacial period. The most detailed evidence comes from the wider area of the tributaries of the lower Elbe River and the Ohře River, which will be used as a model study area in the following text. The obvious starting point for the succession of European rivers floodplains was the last Ice Age. In the prevalently forest-free landscape rivers formed wild streams (Pokorný, 2011) causing strong erosion and transporting inorganic material. In warmer landscapes or during the warmer periods, so called gallery forests formed around the wild streams, for example in the river floodplain of the Slovak part of the Danube River. Apparently, they could have a character of small forests as confirm the presence of a strictly woodland land-snail species climbing on tree trunks of taller trees, Ena montana, in Zlatná u Dunaje (Juřičková et al., 2014). At the beginning of the Holocene, rivers became centres of development and spreading of woodlands through the landscape. The landscape around the rivers started covering by vegetation faster thanks to an amount of nutrients deposited in the river floodplains. Remarkable so called river phenomenon occurred where rivers cut into the hard bedrock with deep canyons creating mosaic of habitats of different orientation and humidity. (Jeník & Slavíková, 1964; Ložek, 1988). This can be observed in the middle stretches of the Vltava River or the Berounka River in the Křivoklátsko or in the Bohemian Karst areas. Agriculture and human settlement concentrated around the rivers especially on nutritious chernozem soils from the Neolithic age. River alluvia were gradually changing as a result of the Neolithic people activity into an agricultural, more or less forest-free landscape (Jiráň & 24 Journal of Landscape Ecology (2015), Vol: 8 / No. 3 Venclová, 2007–2008; Pleinerová & Pavlů, 1979; Pleinerová, 1990) prevented by these activites from spreading of woodlands as well as woodland species of gastropods. Forest-free landscape is in some parts of Bohemia also preserved due to very low precipitation and relatively high average temperatures and is naturally supported by the sub-continental climate as confirmed in palaeobotanical surveys from Zahájí on the lower stream of the Ohře River (Pokorný, 2005; Pokorný et al., 2015) or in the palaeomalacological survey of the neighbouring České Středohoří Mts. (Ložek, 1963; 1964a,b; 1976; 2005; Juřičková et al., 2013a). Climate and human agricultural settlements thus resulted in preserving the mosaic of woodlands, wetlands and open landscape in some areas during the whole Holocene as can be seen in the lower stream of the Ohře River and in the České Středohoří Mts. (Juřičková et al., 2013a,b; Pokorný et al., 2015). The arrival of agriculture caused a kind of return to glacial conditions in that sense that the open landscape deforested by agriculture became a subject to further, intense erosion. Due to the fact that the eroded landscape was before the arrival of agriculture covered by woodland to a large extent, transported soils sedimented as huge alluvial material that became substrate for floodlplain forests. A mosaic of woodland and forest-free areas was formed this way and became typical for the entire area of nutritious soils in the Czech Republic (Ložek, 1964a,b; Pokorný, 2005). Despite the fact it is difficult to confirm this exactly, the up-to-date research of Quaternary sediments in the river floodplains of Czech as well as west-European Rivers suggests that current floodplain forests are younger than Neolithic colonisation and it is related to erosion in landscape caused by the first farmers (Evans, 1993; Lespez et al., 2008; Pišút & Čejka, 2002; Juřičková et al., 2013a,b). This process is not possible to confirm without detailed survey of fossiliferous sediments, because nowadays floodplain forests give the impression of primeval woodlands. As an example can serve the Myslivna Nature Reserve on the Ohře River protected as an original natural old floodplain forest. However, based on fossil mollusc succession (Juřičková et al., 2013a) and composition of recent malacocoenoses that are missing mainly woodland species of gastropods (Horáčková et al., 2014b) we consider this protected floodplain forest as relatively recent. While there are recently living many woodland snail species in the upper stretches of the Ohře River, almost half of them are missing in the lower stream of the river despite the
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